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TFM Mauro Sanna.Pdf (1.537Mb) UNIVERSIDAD DE OVIEDO Vicerrectorado de Internacionalización y Postgrado Máster en Biotecnología Aplicada a la Conservación y Gestión Sostenible de Recursos Vegetales TRABAJO FIN DE MÁSTER DIVERSIDAD, FILOGEOGRAFÍA Y SISTEMÁTICA DE SAXIFRAGA HIRSUTA S. L. Mauro Sanna 22/Julio/2013 Sanna Mauro, 2013. Diversidad, filogeografía y sistemática de Saxifraga hirsuta s. l. 2 Sanna Mauro, 2013. Diversidad, filogeografía y sistemática de Saxifraga hirsuta s. l. ÍNDICE Resumen .............................................................................................................................................. 4 Introducción ........................................................................................................................................ 5 Antecedentes ................................................................................................................................... 5 Planteamiento y objetivos ............................................................................................................... 7 Material y métodos ............................................................................................................................. 8 Muestras estudiadas ....................................................................................................................... 8 Métodos........................................................................................................................................... 9 Extracción de ADN genómico ...................................................................................................... 9 Amplificación de regiones ........................................................................................................... 9 Alineamiento de las secuencias y análisis filogenético ............................................................. 11 Resultados ......................................................................................................................................... 12 Análisis filogenético de las secuencias de ADN nuclear (ITS) ........................................................ 12 Análisis de la diversidad haplotipica de las secuencias rbcL ......................................................... 14 Análisis de la diversidad haplotipica de la región cloroplástica trnL-F .......................................... 16 Conclusiones ...................................................................................................................................... 24 Referencias bibliográficas .................................................................................................................. 25 Anexo I ............................................................................................................................................... 28 3 Sanna Mauro, 2013. Diversidad, filogeografía y sistemática de Saxifraga hirsuta s. l. Resumen: Sanna M. 2013. Diversidad, filogeografía y sistemática de Saxifraga hirsuta s. l. Saxifraga hirsuta subsp. hirsuta L. (Saxifragaceae, sección Gymnopera) es una planta perenne que crece en el norte de la Península Ibérica y en Irlanda, Saxifraga hirsuta subsp. paucicrenata (Gillot) D. A. Webb (Saxifragaceae) es una planta perenne orófila de la cordillera Cantábrica y de los Pirineos. Este estudio se plantea evaluar la diversidad genética de las dos plantas y construir un modelo filogeográfico mediante técnicas moleculares, confrontándolas también con todas las especies de la sección Gymnopera y de otras secciones del genero. ADN nuclear (región ITS) y ADN cloroplástico (rbcL y tnrL-F) han sido utilizados como marcadores genéticos. El análisis de la red de haplotipos y la construcción de arboles filogenéticos ha permitido evaluar que no existe ningún patrón altitudinal en S. hirsuta s. l. y su similitud genética con la especie pirenaica S. umbrosa L. permite considerar las dos especies como una única unidad taxonómica. Otra especie de la misma sección, S. spathularis Brot., forma un robusto clado hermano con S. hirsuta s. l. hecho que permite teorizar sobre la existencia de un ancestro común distribuido geográficamente desde Irlanda hasta la Península Ibérica. La otra especie de la sección, S. cuneifolia L., parece tener un origen más próximo a otras secciones. Este trabajo plantea las bases para un estudio más detallado de toda la sección Gymnopera. Summary: Sanna M. 2013. Diversity, phylogeography and systematic of Saxifraga hirsuta s. l. Saxifraga hirsuta subsp. hirsuta L. (Saxifragaceae, section Gymnopera) is a perennial plant that grows in the North of the Iberian Peninsula and Ireland, Saxifraga hirsuta subsp. paucicrenata (Gillot) D. A. Webb (Saxifragaceae) is a perennial and orophilous plant that lives in the Cantabrian Mountains and the Pyrenees. In this study we suggest the evaluation of the genetic diversity of both species and the construction of phylogeographical models by using molecular techniques as well as the comparison with other species from the section Gymnopera and other sections of the genus. Nuclear DNA (ITS region) and chloroplastic DNA (rbcL and tnrL-F) have been used as genetic markers. The analysis of the haplotype networks as well as the construction of phylogenetic trees has allow us to set that there is not any altitudinal zonation related to S. hirsuta s. l. Moreover, its genetic similarity with the Pyrenean species S. umbrosa L. indicates that both species belong to the same taxonomic unit. The fact that S. spathularis Brot., from the same section performs a strong sister clade with S. hirsuta s. l. allow us to theorize about the existence of a common ancestor geographically distributed from Ireland to the Iberian Peninsula while the other species of the section, S. cuneifolia L., seems to have a more close origin to other sections. This work is the basis for further studies on the whole Gymnopera section. 4 Sanna Mauro, 2013. Diversidad, filogeografía y sistemática de Saxifraga hirsuta s. l. INTRODUCCIÓN Antecedentes El género Saxifraga L. -familia Saxifragaceae, orden Saxifragales (APG 2009)- comprende, si se excluye de él Micranthes Haw., entre 370 y 390 especies distribuidas fundamentalmente por los territorios templados, mediterráneos y árticos del hemisferio norte, aunque algunas crecen en Sudamérica (Brouillet & Elvander 2007; Soltis 2007). El modelo sistemático actual del género distingue unas quince secciones (Gornall 1987; Webb 1987a), entre ellas Saxifraga sect. Gymnopera descrita por Don (1822) incluyendo alguna especie más de las actualmente incluidas en este taxon que, tal como se delimita actualmente, agrupa fundamentalmente plantas europeas occidentales (Webb 1987a, 1993; Vargas 1997): Saxifraga cuneifolia L., Sp. Pl., ed. 2: 574 (1762), que se distribuye desde el Pirineo Oriental hasta los Cárpatos orientales. De acuerdo con Webb (1988, 1993) se distinguen dos razas con categoría subespecífica: S. cuneifolia subsp. cuneifolia y S. cuneifolia subsp. robusta D.A.Webb, Bot. J. Linn. Soc. 97: 355 (1988). Saxifraga hirsuta L., Syst. Nat., ed. 10, 2: 1026 (1759) designada por Gornall (1987) tipo de la sección y distribuida desde la cordillera Cantábrica hasta los Pirineos centrales e Irlanda. Se distinguen en su seno dos subespecies: S. hirsuta subsp. hirsuta y S. hirsuta subsp. paucicrenata (Gillot) D.A.Webb, Feddes Repert. 68: 201 (1963), esta última una raza orófila de la cordillera Cantabrica y los Pirineos (Webb 1993; Vargas 1997). Saxifraga spathularis Brot., Fl. Lusit. 2: 172 (1805), de distribución ibérica noroccidental e hibérnica. Saxifraga umbrosa L., Sp. Pl., ed. 2: 574 (1762), distribuida por los Pirineos occidentales y centrales. Además, se han descrito tres híbridos entre especies de esta sección: Saxifraga ×polita (Haw.) Link, Enum. Hort. Berol. Alt. 1: 414 (1821) (S. hirsuta x S. spathularis), S ×geum L., Sp. Pl.: 401 (1753) (S. hirsuta x S. umbrosa) y S. ×urbium D.A.Webb, Feddes Repert. 68: 199 (1963) (S. spathularis x S. umbrosa). La sistemática tradicional del género (Haworth 1812, 1821; Don 1822; Engler & Irmscher, 1919) ha sufrido escasos cambios, salvo la ya indicada segregación de Micranthes Haw. que ha sido consecuencia, fundamentalmente, de las conclusiones derivadas del uso de marcadores moleculares (Soltis et al. 1996, 2001; Fernández 5 Sanna Mauro, 2013. Diversidad, filogeografía y sistemática de Saxifraga hirsuta s. l. Prieto et al. 2013). Estos criterios sistemáticos, apoyados en el análisis de secuencias de ADN tanto nuclear como cloroplástico (Soltis et al. 1996, 2001; Conti et al. 1999; Vargas et al. 1999; Vargas 2000), han permitido confirmar, en la mayoría de los casos, la sistemática seccional establecida básicamente por Gornall (1987) y comúnmente aceptada (Webb 1987a, 1993; Vargas 1997; Gornall et al. 2000; Jintang et al. 2001; McGregor 2008). El estudio de Saxifraga sect. Gymnopera mediante marcadores moleculares se ha desarrollado escasamente, aunque secuencias de muestras de especies de esta sección han sido incluidas en estudios de otros grupos del género; el análisis de secuencias ITS de muestras de S. cuneifolia, S. hirsuta y S. spathularis ha llevado a Conti et al. (1999) a concluir que Saxifraga sect. Gymnopera constituye un clado monofilético. Pérez Vallines (2011) investigó, mediante el análisis secuencias de ADN tanto nuclear (ITS) como cloroplástico (rbcL), el origen y diversidad de las poblaciones de S. hirsuta y S. spathularis de la Península Ibérica e Irlanda, evidenciándose la posibilidad de usar esas u otras secuencias para estudiar
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